Malaysia, Sabah, Ranau, Gunung Kinabalu, 6.0428°N, 116.5587°E, 2600 m.

Malaysia: Holotype ♂, Sabah: Ranau, Gunung Kinabalu, 6.0428°N, 116.5587°E, 2600 m, 2.v.1987, Syzygium korthalsianum (D. Burckhardt & I. Löbl) #8751 (MNHG, slide mounted). – Malaysia: 6 ♂, 8 ♀, same but summit trail, 3230 m, Leptospermum sp., 29.iv.1982, Leptospermum scrub (D. Burckhardt) #8277; 10 ♂, 12 ♀, same but 3230 m, Leptospermum recurvum, #8278; 1 ♂, 1 ♀, same but 2600 m, 1.v.1987 (D. Burckhardt & I. Löbl) #8735; 21 ♂, 36 ♀, same but 2600 m, 2.v.1987, Leptospermum recurvum, #8747, #8748, #8749; 6 ♂, 6 ♀, same but 2600 m, 2.v.1987, general sweeping of vegetation, #8752; 2 ♂, 9 ♀, same but 2600 m, 2.v.1987, Leptospermum javanicum, #8753; 12 ♂, 13 ♀, same but 3300 m, 4.v.1987, Leptospermum recurvum, #8756; 4 ♂, 4 ♀, same but below Layang Layang, 2600 m, 2‒8.v.1987, interception trap (A. Smetana); 7 ♂, 11 ♀, same but below Laban Rata, 3155 m, 5.v.1987; 1 ♂, 5 ♀, same but Laban Rata, 3200 m, 4‒8.v.1987, interception trap; 8 ♂, 7 ♀, same but 3200 m, 9‒20.v.1987, interception trap (MHNG, NHMB, dry and slide mounted).

Genal processes 0.3 times as long as vertex along mid-line, irregularly rounded anteriorly. Forewing oblong oval, widest in the middle, 2.3–2.7 times as long as broad, broadly rounded apically; vein C+Sc mostly straight, weakly concave in proximal third, cell c+sc narrow. Surface spinules present in all cells, forming cellular pattern; in cell r₂ above bifurcation of vein M, the cells are irregularly hexagonal consisting of two indistinct rows of spinules. No extra pore fields developed on abdominal intersegmental membrane. Basal segment of proctiger weakly curved posteriorly, hind margin with a row of stout setae; apical segment 0.2–0.3 times as long as basal segment. Paramere digitiform. Distal segment of aedeagus cuneate. Female terminalia strongly narrowed medially, bearing each an apical process on proctiger and subgenital plate, female proctiger dorsally serrate.

Adult. Colouration. Head and thorax light reddish brown. Vertex with dark brown dot in the middle of either half; genal processes dark brown at base, yellow apically. Antenna light orange brown at base, getting gradually darker from segment 6 to apex which is dark brown or black. Pronotum with each two submedian dark dots and mesopraescutum with each one submedian dark dot along fore margin on either side. Legs yellow; profemora light greyish brown. Forewings light ochreous or amber-coloured, slightly lighter along fore margin; veins concolourous with membrane. Abdomen yellow or orange; base of female proctiger light brown. Younger specimens with less extended dark colour.

Conforming to the generic description of Burckhardt et al. (2020). Body length ♂ 1.4–1.5 mm, ♀ 1.5–1.9 mm (6 ♂, 6 ♀). Head deflexed 45° from longitudinal axis of body. Vertex rhomboidal, weakly concave at base; preocular sclerite forming flat tubercule; genal processes 0.3 times as long as vertex along mid-line, irregularly rounded anteriorly, contiguous medially; eyes weakly ‘stalked’ (Fig. 1). Antenna 0.6–1.0 times as long as head width. Metatibia 0.5–0.7 times as long as head width, weakly widening to apex, with 5 irregularly spaced apical spurs. Forewing (Fig. 4) oblong oval, widest in the middle, 2.3–3.0 times as long as head width, 2.3–2.7 times as long as broad, broadly rounded apically; pterostigma, at base narrower than adjacent part of cell r₁, regularly narrowing to apex, ending at apical fifth of wing; vein C+Sc mostly straight, weakly concave in proximal third, cell c+sc narrow; vein Rs almost straight, vein M long, with short, weakly diverging branches, vein Cu_1a relatively straight, reaching the wing margin distinctly distal to bifurcation of vein M. Surface spinules present in all cells, forming cellular pattern; in cell r₂ above bifurcation of vein M, the cells are irregularly hexagonal consisting of two indistinct rows of spinules (Fig. 5). No extra pore fields developed on abdominal intersegmental membrane. Male terminalia as in Figs 10–12. Proctiger 0.4–0.6 times as long as head width; basal segment, in profile, weakly curved posteriorly, irregularly beset with fine setae, hind margin with a row of stout setae; apical segment tubular, 0.2–0.3 times as long as basal segment. Subgenital plate, in profile, triangular, with almost straight dorsal margin; sparsely beset with short setae. Paramere, in profile, digitiform, almost straight anteriorly, wavy posteriorly, broadly rounded apically; inner face densely coverd in moderately long bristles. Distal portion of aedeagus slender in basal half, widening towards apex which is rounded; sclerotised end tube of ductus ejaculatorius small, weakly curved. Female terminalia as in Fig. 13. Proctiger 0.8–1.0 times as long as head width, 2.3–3.7 times as long as circumanal ring, cuneate; strongly narrowed medially, bearing an apical process, apical third serrate dorsally, subacute apically; sparsely beset with short setae, with a longitudinal lateral row of slightly longer setae in apical third and 2 longitudinal rows of peg setae near ventral margin in apical half. Subgenital plate 0.7–0.8 times as long as proctiger, in profile, bearing narrow, apically pointed process. Valvulae dorsalis and ventralis straight (Fig. 14); valvula lateralis pointed apically.

Measurements in mm (5 ♂, 6 ♀). Head width 0.44–0.58; antenna length 0.34–0.50; forewing length 1.08–1.48; length of male proctiger 0.20–0.24; paramere length 0.16–0.18; length of distal portion of aedeagus 0.10–0.14; female proctiger length 0.42–046.

Fifth instar immature unknown.

Malaysia: Sabah, probably endemic to Gunung Kinabalu.

Adults were collected in large numbers on Leptospermum javanicum Blume (= L. flavescens auct.) and L. recurvum Hook.f. (Myrtaceae) suggesting that these two species constitute hosts. Leptospermum javanicum is widely distributed from Burma and southern Thailand to the Philippines, Malucu and Lesser Sunda Islands, and L. recurvum, is endemic to Gunung Kinabalu (Thompson 1989). It has been suggested that L. recurvum has split from the former after the last Pleistocene glaciation (Lee and Lowry 1980). The species occurs as a tree and at high altitudes as prostrate shrub on an outcrop of ultra basic rocks. It is one of the main shrub species of the summit zone above 3200 m altitude (Cockburn 1978; Corner 1978). A singe male was collected also on Syzygium korthalsianum (Miq.) Miq. (Myrtaceae), which is an unlikely host contrary to the statement by Burckhardt et al. (2020).

Ctenarytaina daleae is most similar to C. insularis Martoni & Armstrong in the posteriorly weakly lobed male proctiger, the digitiform paramere, the female terminalia, which are strongly narrowed medially and bear each an apical process on the proctiger and subgenital plate as well as the dorsally serrate female proctiger. It differs from the latter in the surface spinules forming rings consisting of two rows of spinules, the distal aedeagal segment which is evenly widening to apex rather than with a slender stalk and inflated apical part, the dorsally less concave female proctiger and the host plant: Leptospermum versus Syzygium.

Ctenarytaina daleae was described based on the male holotype and two female paratypes (Burckhardt et al. 2020). The examination of a long series of material from the summit region of Gunung Kinabalu collected on Leptospermum spp. shows that the holotype of C. daleae corresponds to the species from Leptospermum but not the females which belong to C. smetanai sp. nov. described below. Ctenarytaina daleae and C. smetanai sp. nov. are similar in the head and forewing structure (Figs 1, 3, 4, 8) but differ in the male and female terminalia (Figs 10–14, 18–20, 23, 24).

Figures 1–3. 

Ctenarytaina spp., head, in dorsal view. 1. C. daleae; 2. C. lienhardi sp. nov.; 3. C. smetanai sp. nov.

Figures 4–9. 

Ctenarytaina spp. 4, 6, 8. Forewing; 5, 7, 9. Surface spinules in cell r₂ above bifurcation of vein M. 4, 5. C. daleae; 6, 7. C. lienhardi sp. nov.; 8, 9. C. smetanai sp. nov.

Malaysia, Sabah, Ranau, Gunung Kinabalu, 1750–1800 m.

Holotype ♂, Malaysia: Sabah: Ranau, Gunung Kinabalu, 1750–1800 m, 22.iii.1983 (C. Lienhard) #Pal-83/47 (MHNG, dry mounted).

Paratypes : Malaysia: 1 ♂, 2 ♀, same data as holotype; 1 ♀, same but, 1750 m, 27.iv.1987, Tristaniopsis (D. Burckhardt and I. Löbl) #8722; 1 ♀, same but, 1750–1850 m, 20.iii.1983 (C. Lienhard) #Pal-83/34 (MHNG, NHMB, dry and slide mounted).

Genal processes 0.3 times as long as vertex along mid-line, irregularly rounded anteriorly. Forewing oblong oval, widest in the middle, 2.3–2.5 times as long as broad, narrowly rounded apically; vein C+Sc mostly straight, cell c+sc narrow. Surface spinules present in all cells, forming cellular pattern; in cell r₂ above bifurcation of vein M, the cells are irregularly hexagonal consisting of one or two indistinct rows of spinules. Extra pore fields developed on abdominal intersegmental membrane. Basal segment of proctiger weakly curved posteriorly, hind margin with a row of stout setae; apical segment 0.2 times as long as basal segment. Paramere weakly cuneate; inner face densely beset with long bristles. Distal portion of aedeagus slender in basal half, weakly inflated in apical half. Female terminalia cuneate; proctiger blade-shaped distal to circumanal ring, smooth dorsally, blunt apically.

Adult. Colouration. Head and thorax light reddish brown. Vertex with dark brown dot in the middle of either half; genal processes dark brown at base, yellow apically. Antenna light orange brown at base, getting gradually darker from segment 6 to apex which is dark brown or black. Pronotum with each two submedian dark dots and mesopraescutum with each one submedian dark dot along fore margin on either side. Legs yellow; profemora light greyish brown. Forewings light ochreous or amber-coloured, slightly lighter along fore margin; veins concolourous with membrane. Abdomen yellow or orange; base of female proctiger light brown. Younger specimens with less extended dark colour.

Conforming to the generic description of Burckhardt et al. (2020). Body length ♂ 2.0 mm, ♀ 2.4 mm (1 ♂, 1 ♀). Head deflexed 45° from longitudinal axis of body. Vertex rhomboidal, weakly concave at base; preocular sclerite forming small tubercule; genal processes 0.3 times as long as vertex along mid-line, irregularly rounded anteriorly, contiguous medially; eyes weakly ‘stalked’ (Fig. 2). Antenna 0.8 times as long as head width. Metatibia 0.5–0.6 times as long as head width, weakly widening to apex, with 5 irregularly spaced apical spurs. Forewing (Fig. 6) oblong oval, widest in the middle, 2.3–2.5 times as long as head width, 2.5–2.6 times as long as broad, narrowly rounded apically; pterostigma, at base wider than adjacent part of cell r₁, regularly narrowing to apex, ending at apical quarter of wing; vein C+Sc mostly straight, cell c+sc narrow; vein Rs almost straight, vein M long, with short, weakly diverging branches, vein Cu_1a relatively straight, reaching the wing margin at bifurcation of vein M. Surface spinules present in all cells, forming cellular pattern; in cell r₂ above bifurcation of vein M, the cells are irregularly hexagonal consisting of one or two rows of surface spinules (Fig. 7). Extra pore fields present on abdominal intersegmental membrane. Male terminalia as in Figs 15–17. Proctiger 0.5 times as long as head width; basal segment, in profile, weakly curved posteriorly; irregularly beset with fine setae, bearing a row of short bristles along hind margin; apical segment tubular, 0.2 times as long as basal segment. Subgenital plate, in profile, triangular, slightly elongate, with weakly concave dorsal margin; sparsely beset with short setae. Paramere, in profile, weakly cuneate, relatively straight, narrowly rounded apically; inner face densely beset with long bristles. Distal portion of aedeagus slender in basal half, weakly inflated in apical half; sclerotised end tube of ductus ejaculatorius small, almost straight. Female terminalia as in Fig. 21. Proctiger 1.0 times as long as head width, 2.7–2.8 times as long as circumanal ring, cuneate; dorsal margin of proctiger concave, blade-shaped distal to circumanal ring, smooth dorsally, blunt apically; sparsely beset with short setae, with a longitudinal lateral row of slightly longer setae in apical half and 3 longitudinal rows of peg setae near ventral margin in apical half. Subgenital plate 0.7–0.8 times as long as proctiger, in profile, forming narrow process in apical half, pointed apically. Valvulae dorsalis and ventralis weakly curved (Fig. 22); valvula lateralis narrowly rounded apically.

Measurements in mm (1 ♂, 2 ♀). Head width 0.64–0.74; antenna length 0.60; forewing length 1.54–1.84; length of male proctiger 0.38; paramere length 0.28; length of distal portion of aedeagus 0.16; female proctiger length 0.64–0.72.

Fifth instar immature unknown.

Named after C. Lienhard, Genève, eminent specialist of Psocodea and collector of most of the type series.

Malaysia: Sabah, probably endemic to Mount Kinabalu.

Unknown; a single female was collected on Tristaniopsis (Myrtaceae).

Ctenarytaina lienhardi sp. nov. shares with C. daleae, C. insularis and C. taylori the narrow forewings and the posteriorly weakly curved male proctiger. It differs from the two former species in the blade-shaped, dorsally smooth female proctiger and from the latter in the lack of a dark brown basal patch on the forewing.

Malaysia, Sabah, Ranau, Gunung Kinabalu, below Layang Layang, 2600 m.

Holotype ♂. Malaysia: Sabah: Ranau, Gunung Kinabalu, below Layang Layang, 2600 m, 2–8.v.1987, interception trap (A. Smetana) (MHNG, slide mounted).

Paratypes. Malaysia, 1 ♀, Sabah, Ranau, Gunung Kinabalu, same as holotype (dry mounted); 1 ♀, same but 2600 m, 1.v.1987, Syzygium punctilimbum (D. Burckhardt & I. Löbl) #8737 (MNHG, slide mounted); 1 ♀, same but Gunung Kinabalu, 6.0195°N, 116.5385°E, 1700 m, 22.iii.1983 (S. Nagai) #Pal-83/46 (MHNG, dry mounted).

Genal processes 0.5 times as long as vertex along mid-line, irregularly rounded anteriorly. Forewing oblong oval, widest in the middle, 1.9–2.8 times as long as broad, narrowly rounded apically; vein C+Sc mostly straight, weakly concave in proximal third, cell c+sc narrow. Surface spinules present in all cells, forming cellular pattern; in cell r₂ above bifurcation of vein M, the cells are irregularly hexagonal consisting of one row of spinules. No extra pore fields developed on abdominal intersegmental membrane. Basal segment trapezoidal, hind margin with subapical angle bearing 5 short peg setae; apical segment 0.2–0.3 times as long as basal segment. Paramere very long and slender. Distal portion of aedeagus slender, weakly inflated apically. Female terminalia cuneate; dorsal margin of proctiger angularly concave at caudal end of circumanal ring, from there to pointed apex weakly convex, smooth.

Adult. Colouration. Head and thorax dark reddish brown to almost black. Genal processes lighter than vertex, ochreous to brown. Antennal segments 1 and 2 light brown, segments 3–5 and basal third of segment 6 yellow to ochreous, apical two thirds of segment 6 and segments 7–10 dark brown to almost black. Mesoscutum with a broader median and two narrower submedian reddish brown longitudinal stripes. Fore and mid legs brown, tarsi and hind legs, including metacoxae, dirty yellowish. Forewings amber-coloured or light brown; veins concolourous with membrane. Hindwings whitish. Abdomen reddish brown to almost black dorsally, black ventrally; male and female terminalia light brown.

Conforming to the generic description of Burckhardt et al. (2020). Body length ♂ 1.9 mm, ♀ 2.0–2.2 mm (1 ♂, 2 ♀). Head deflexed 45° from longitudinal axis of body. Vertex rhomboidal, concave at base; preocular sclerite forming small tubercule; genal processes 0.5 times as long as vertex along mid-line, irregularly rounded anteriorly, well separated medially; eyes moderately ‘stalked’ (Fig. 3). Antenna 0.9–1.1 times as long as head width. Metatibia 0.5 times as long as head width, weakly widening to apex, with 5 irregularly spaced apical spurs. Forewing (Fig. 8) oblong oval, widest in the middle, 2.3–2.4 times as long as head width, 1.9–2.8 times as long as broad, narrowly rounded apically; pterostigma, at base narrower than adjacent part of cell r₁, regularly narrowing to apex, ending at apical third of wing; vein C+Sc mostly straight, weakly concave in proximal third, cell c+sc narrow; vein Rs almost straight, vein M long, with short, weakly diverging branches, vein Cu_1a relatively straight, reaching the wing margin distinctly distal to bifurcation of vein M. Surface spinules present in all cells, forming cellular pattern; in cell r₂ above bifurcation of vein M, the cells are irregularly hexagonal consisting of one row of spinules (Fig. 9). No extra pore fields developed on abdominal intersegmental membrane. Male terminalia as in Figs 18–20. Proctiger 0.7 times as long as head width; basal segment, in profile, trapezoidal; irregularly beset with setae along anterior and posterior margins, hind margin with subapical angle bearing 5 short peg setae; apical segment tubular, 0.3 times as long as basal segment. Subgenital plate, in profile, elongate, with wavy dorsal margin; sparsely beset with short setae. Paramere, in profile, very long and slender, slightly narrowed in apical third, narrowly rounded apically; inner face with moderately long bristles in apical third and with two rows of bristles in basal two thirds. Distal portion of aedeagus slender, weakly inflated apically; sclerotised end tube of ductus ejaculatorius small, relatively straight. Female terminalia as in Fig. 23. Proctiger 1.1 times as long as head width, 2.9 times as long as circumanal ring, cuneate; dorsal margin of proctiger angularly concave at caudal end of circumanal ring, from there to pointed apex weakly convex, smooth; weakly beset with short setae, with a longitudinal lateral row of long setae in apical third and 2–3 longitudinal rows of peg setae near ventral margin in apical half. Subgenital plate 0.7 times as long as proctiger, in profile, evenly narrowing to pointed apex. Valvulae dorsalis and ventralis curved (Fig. 24).

Figures 10–24. 

Ctenarytaina spp., terminalia. 10, 15, 18. Male terminalia, in profile; 11, 16, 19. Distal portion of aedeagus; 12, 17, 20. Inner face of paramere; 13, 21, 23. Female terminalia, in profile; 14, 22, 24. Dorsal and ventral valvulae. 10–14 C. daleae; 15–17, 21, 22. C. lienhardi sp. nov.; 18–20, 23, 24. C. smetanai sp. nov.

Measurements in mm (1 ♂, 1 ♀). Head width 0.62–0.72; antenna length 0.64–0.66; forewing length 1.46–1.68; length of male proctiger 0.46; paramere length 0.44; length of distal portion of aedeagus 0.20; female proctiger length 0.82.

Fifth instar immature unknown.

Named after the late A. Smetana, Ottawa, eminent staphylinid specialist and collector of the holotype.

Malaysia: Sabah, probably endemic to Mount Kinabalu.

One female was swept from Syzygium punctilimbum (Merr.) Merr. & L.M.Perry (Myrtaceae), a possible host, in moss forest. One male and one female were collected in an interception trap and one female was taken by sweeping vegetation.

Ctenarytaina smetanai sp. nov. differs from other congeners in the basal segment of the male proctiger which is trapezoidal and bears 5 short peg setae on the subapical angle of hind margin (Fig. 18).